Photosynthetic, hydraulic and biomechanical responses of Juglans californica shoots to wildfire

Leaf gas exchange and stem xylem hydraulic and mechanical properties were studied for unburned adults and resprouting burned Juglans californica (southern California black walnut) trees 1 year after a fire to explore possible trade-offs between mechanical and hydraulic properties of plants. The CO₂ uptake rates and stomatal conductance were 2–3 times greater for resprouting trees than for unburned adults. Both predawn and midday water potentials were more negative for unburned adult trees, indicating that the stems were experiencing greater water stress than the stems of resprouting trees. In addition, the xylem specific conductivity was similar in the two growth forms, even though the stems of resprouting trees were less vulnerable to water-stress-induced embolism than similar diameter, but older, stems of adult trees. The reduced vulnerability may have been due to less cavitation fatigue in stems of resprouts. The modulus of elasticity, modulus of rupture and xylem density were all greater for resprouts, indicating that resprouts have greater mechanical strength than do adult trees. The data suggest that there is no trade-off between stem mechanical strength and shoot hydraulic and photosynthetic efficiency in resprouts, which may have implications for the success of this species in the fire-prone plant communities of southern California.     

Neither xylem collapse,cavitation, or changing leaf conductance drive stomatal closure in wheat

Identifying the drivers of stomatal closure and leaf damage during stress in grasses is a critical prerequisite for understanding crop resilience. Here, we investigated whether changes in stomatal conductance (g_s) during dehydration were associated with changes in leaf hydraulic conductance (K_leaf), xylem cavitation, xylem collapse, and leaf cell turgor in wheat (Triticum aestivum). During soil dehydration, the decline of g_s was concomitant with declining K_leaf under mild water stress. This early decline of leaf hydraulic conductance was not driven by cavitation, as the first cavitation events in leaf and stem were detected well after K_leaf had declined. Xylem vessel deformation could only account for <5% of the observed decline in leaf hydraulic conductance during dehydration. Thus, we concluded that changes in the hydraulic conductance of tissues outside the xylem were responsible for the majority of K_leaf decline during leaf dehydration in wheat. However, the contribution of leaf resistance to whole plant resistance was less than other tissues (<35% of whole plant resistance), and this proportion remained constant as plants dehydrated, indicating that K_leaf decline during water stress was not a major driver of stomatal closure.     

Wood structure and function change with maturity: Age of the vascular cambium is associated with xylem changes in current‐year growth

Xylem vessel structure changes as trees grow and mature. Age‐ and development‐related changes in xylem structure are likely related to changes in hydraulic function. We examined whether hydraulic function, including hydraulic conductivity and vulnerability to water‐stress‐induced xylem embolism, changed over the course of cambial development in the stems of 17 tree species. We compared current‐year growth of young (1–4 years), intermediate (2–7 years), and older (3–10 years) stems occurring in series along branches. Diffuse and ring porous species were examined, but nearly all species produced only diffuse porous xylem in the distal branches that were examined irrespective of their mature xylem porosity type. Vessel diameter and length increased with cambial age. Xylem became both more conductive and more cavitation resistant with cambial age. Ring porous species had longer and wider vessels and xylem that had higher conductivity and was more vulnerable to cavitation; however, these differences between porosity types were not present in young stem samples. Understanding plant hydraulic function and architecture requires the sampling of multiple‐aged tissues because plants may vary considerably in their xylem structural and functional traits throughout the plant body, even over relatively short distances and closely aged tissues.     

Photosynthetic pathway alters xylem structure and hydraulic function in herbaceous plants

Plants using the C₄ photosynthetic pathway have greater water use efficiency (WUE) than C₃ plants of similar ecological function. Consequently, for equivalent rates of photosynthesis in identical climates, C₄ plants do not need to acquire and transport as much water as C₃ species. Because the structure of xylem tissue reflects hydraulic demand by the leaf canopy, a reduction in water transport requirements due to C₄ photosynthesis should affect the evolution of xylem characteristics in C₄ plants. In a comparison of stem hydraulic conductivity and vascular anatomy between eight C₃ and eight C₄ herbaceous species, C₄ plants had lower hydraulic conductivity per unit leaf area (K_L) than C₃ species of similar life form. When averages from all the species were pooled together, the mean K_L for the C₄ species was 1.60 × 10^?4 kg m^?1 s^?1 MPa^?1, which was only one‐third of the mean K_L of 4.65 × 10^?4 kg m^?1 s^?1 MPa^?1 determined for the C₃ species. The differences in K_L between C₃ and C₄ species corresponded to the two‐ to three‐fold differences in WUE observed between C₃ and C₄ plants. In the C₄ species from arid regions, the difference in K_L was associated with a lower hydraulic conductivity per xylem area, smaller and shorter vessels, and less vulnerable xylem to cavitation, indicating the C₄ species had evolved safer xylem than the C₃ species. In the plants from resource‐rich areas, such as the C₄ weed Amaranthus retroflexus, hydraulic conductivity per xylem area and xylem anatomy were similar to that of the C₃ species, but the C₄ plants had greater leaf area per xylem area. The results indicate the WUE advantage of C₄ photosynthesis allows for greater flexibility in hydraulic design and potential fitness. In resource‐rich environments in which competition is high, an existing hydraulic design can support greater leaf area, allowing for higher carbon gain, growth and competitive potential. In arid regions, C₄ plants evolved safer xylem, which can increase survival and performance during drought events.     

Beating the blaze: Fire survival in the fan aloe (Kumara plicatilis), a succulent monocotyledonous tree endemic to the Cape fynbos,South Africa

Fire is central to the ecology of Mediterranean‐type climate ecosystems, but little is known about the fire ecology of succulent plants therein. This study investigated the fire ecology of an arborescent succulent monocot, Kumara plicatilis (L.) G. D. Rowley (Asphodelaceae), a Cape fynbos endemic. Habitat suitability was assessed to determine whether the species tolerates or ‘avoids’ fire, and fire survival traits (bark thickness and tissue water content) were measured. The population size structure and density of three K. plicatilis populations were assessed after natural fires, and resprouting potential was investigated. Kumara plicatilis adopts a dual fire survival strategy, occupying rocky sites to ‘avoid’ fire and possessing morphological features that afford fire tolerance, e.g. well‐protected apical meristems and thick corky bark. Bark thickness of burned individuals in situ was similar to unburned plants, suggesting that K. plicatilis bark provides effective insulation against fire. Mortality rates were 64%, 40% and 11%, and decreased as rock cover at the population level increased. All three populations showed reduced plant density post‐fire, with greater density reductions associated with lower rock cover. Small plants appear most vulnerable to fire damage due to lower absolute bark thickness and plant heights within the flame zone. Kumara plicatilis is an apical sprouter, recovering after fire or mechanical stem damage by onward growth from surviving stem apices, rather than resprouting. Post‐fire population recovery therefore likely depends on inter‐fire recruitment.     

The physiological implications of primary xylem organization in two ferns

Xylem structure and function are well described in woody plants, but the implications of xylem organization in less‐derived plants such as ferns are poorly understood. Here, two ferns with contrasting phenology and xylem organization were selected to investigate how xylem dysfunction affects hydraulic conductivity and stomatal conductance (g_s). The drought‐deciduous pioneer species, Pteridium aquilinum, exhibits fronds composed of 25 to 37 highly integrated vascular bundles with many connections, high g_s and moderate cavitation resistance (P₅₀ = ?2.23 MPa). By contrast, the evergreen Woodwardia fimbriata exhibits sectored fronds with 3 to 5 vascular bundles and infrequent connections, low g_s and high resistance to cavitation (P₅₀ = ?5.21 MPa). Xylem‐specific conductivity was significantly higher in P. aqulinium in part due to its wide, efficient conduits that supply its rapidly transpiring pinnae. These trade‐offs imply that the contrasting xylem organization of these ferns mirrors their divergent life history strategies. Greater hydraulic connectivity and g_s promote rapid seasonal growth, but come with the risk of increased vulnerability to cavitation in P. aquilinum, while the conservative xylem organization of W. fimbriata leads to slower growth but greater drought tolerance and frond longevity.     

Functional lifespans of xylem vessels: Development,hydraulic function,and post‐function of vessels in several species of woody plants

Premise of the Study

Xylem vessels transition through different stages during their functional lifespan, including expansion and development of vessel elements, transition to vessel hydraulic functionality, and eventual transition to post‐functionality. We used information on vessel development and function to develop a model of vessel lifespan for woody plants.

Methods

We examined vessel functional lifespan using repeated anatomical sampling throughout the growing season, combined with active‐xylem staining to evaluate vessel hydraulic transport functionality. These data were combined with a literature review. The transitions between vessel functional lifespans for several species are illustrated, including grapevine (Vitis vinifera L., Vitaceae), English oak (Quercus robur L., Fagaceae), American chestnut [Castanea dentata (Marshall) Borkh.; Fagaceae], and several arid and semi‐arid shrub species.

Key Results

In intact woody plants, development and maturation of vessel elements may be gradual. Once hydraulically functional, vessel elements connect to form a vessel network that is responsible for bulk hydraulic flow through the xylem. Vessels become nonfunctional due to the formation of gas emboli. In some species and under some conditions, vessel functionality of embolized conduits may be restored through refilling. Blockages, such as tyloses, gels, or gums, indicate permanent losses in hydraulic functional capacity; however, there may be some interesting exceptions to permanent loss of functionality for gel‐based blockages.

Conclusions

The gradual development and maturation of vessel elements in woody plants, variation in the onset of functionality between different populations of vessels throughout the growing season, and differences in the timing of vessel transitions to post‐functionality are important aspects of plant hydraulic function.     

The impact of vessel size on vulnerability curves: data and models for within‐species variability in saplings of aspen,Populus tremuloides Michx

The objective of this study was to quantify the relationship between vulnerability to cavitation and vessel diameter within a species. We measured vulnerability curves (VCs: percentage loss hydraulic conductivity versus tension) in aspen stems and measured vessel‐size distributions. Measurements were done on seed‐grown, 4‐month‐old aspen (Populus tremuloides Michx) grown in a greenhouse. VCs of stem segments were measured using a centrifuge technique and by a staining technique that allowed a VC to be constructed based on vessel diameter size‐classes (D). Vessel‐based VCs were also fitted to Weibull cumulative distribution functions (CDF), which provided best‐fit values of Weibull CDF constants (c and b) and P₅₀ = the tension causing 50% loss of hydraulic conductivity. We show that P₅₀ = 6.166D^?0.3134 (R² = 0.995) and that b and 1/c are both linear functions of D with R² > 0.95. The results are discussed in terms of models of VCs based on vessel D size‐classes and in terms of concepts such as the ‘pit area hypothesis’ and vessel pathway redundancy.     

Tradeoffs between hydraulic efficiency and mechanical strength in the stems of four co-occurring species of chaparral shrubs

Possible tradeoffs between efficiency of water transport and mechanical strength were examined in stems of two congeneric pairs of co-occurring chaparral shrubs. First, since previously published results indicated that Adenostoma sparsifolium (Rosaceae) had greater specific conductivity (k _s or hydraulic conductivity per xylem transverse area) than A. fasciculatum, it was hypothesized that A. sparsifolium would have greater vessel lumen area per square millimeter of xylem area, and less mechanical strength, than A. fasciculatum. Secondly, since Ceanothus megacarpus (Rhamnaceae) is a non-sprouter (unable to sprout from the root crown following fire or other major disturbance) whereas C. spinosus is a sprouter and thus able to form new stems following disturbance, it was hypothesized that C. megacarpus would have greater mechanical strength, but lower k _s, than C. spinosus. Both hypotheses were supported. Based upon computer-aided image analyses, A. sparsifolum had significantly higher mean and maximum vessel diameters (16.4, 40.5 vs. 14.6, 35.7 μm), a 34% greater percent vessel lumen area, and a two-fold greater measured and theoretical k _s than A. fasciculatum. This corresponded to 14% lower stem density (wet weight/volume) and less mechanical strength, with a 37% lower modulus of elasticity (MOE) and a 30% lower modulus of rupture (MOR) than A. fasciculatum. Similarly, C.␣spinosus had a significantly higher maximum vessel diameter (52.7 vs. 41.8 μm) and a 92% higher theoretical k _s (and 43% higher measured k _s) than C. megacarpus. This corresponded to a 9% lower stem density and 20% lower MOR than for C. megacarpus. Thus, at least within these two congeneric pairs of chaparral shrubs growing together in the same habitat, there may be tradeoffs between mechanical strength and conductive efficiency of the stem xylem which correspond to differences in transport physiology and life history traits of sprouter versus non-sprouter species.     

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought‐induced mortality in Eucalyptus radiata seedlings

It has been reported that elevated temperature accelerates the time‐to‐mortality in plants exposed to prolonged drought, while elevated [CO₂] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO₂] and temperature on the inter‐dependent carbon and hydraulic characteristics associated with drought‐induced mortality in Eucalyptus radiata seedlings grown in two [CO₂] (400 and 640 μL L^?1) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO₂] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO₂], may be the primary contributors to drought‐induced seedling mortality under future climates.     

Common trade‐offs between xylem resistance to cavitation and other physiological traits do not hold among unrelated Populus deltoides ×Populus nigra hybrids

We examined the relationships between xylem resistance to cavitation and 16 structural and functional traits across eight unrelated Populus deltoides×Populus nigra genotypes grown under two contrasting water regimes. The xylem water potential inducing 50% loss of hydraulic conductance (Ψ₅₀) varied from ?1.60 to ?2.40 MPa. Drought‐acclimated trees displayed a safer xylem, although the extent of the response was largely genotype dependant, with Ψ₅₀ being decreased by as far as 0.60 MPa. At the tissue level, there was no clear relationship between xylem safety and either xylem water transport efficiency or xylem biomechanics; the only structural trait to be strongly associated with Ψ₅₀ was the double vessel wall thickness, genotypes exhibiting a thicker double wall being more resistant. At the leaf level, increased cavitation resistance was associated with decreased stomatal conductance, while no relationship could be identified with traits associated with carbon uptake or bulk leaf carbon isotope discrimination, a surrogate of intrinsic water‐use efficiency. At the whole‐plant level, increased safety was associated with higher shoot growth potential under well‐irrigated regime only. We conclude that common trade‐offs between xylem resistance to cavitation and other physiological traits that are observed across species may not necessarily hold true at narrower scales.     

Does acclimation in cavitation resistance due to mechanical perturbation support the pit area or conduit reinforcement hypotheses in Phaseolus vulgaris?

Two Phaseolus vulgaris L. cultivars were exposed to reduced water and stem mechanical perturbation treatments (flexing) to determine if acclimation to these treatments induced hydraulic changes, altered cavitation resistance and changed stem mechanical properties. Additionally, this study sought to determine if changes in cavitation resistance would support the pit area or conduit reinforcement hypotheses. Flexing reduced biomass, leaf area, xylem vessel area and hydraulic conductivity. One cultivar had greater measures of stem strength and cavitation resistance. Flexing increased cavitation resistance (P₅₀) but did not increase Young's modulus, rigidity or flexural strength on dried stems. Stem rigidity and basal diameter were correlated with leaf mass. The ratio of conduit wall thickness to span [(t/b)_h²] increased under high water and flexing treatments while rigidity decreased for one cultivar exposed to both flexing and lower water suggesting an inability to compensate for two simultaneous stresses. Although P₅₀ was not correlated with measures of mechanical strength, P₅₀ was correlated with vessel diameter, consistent with the pit area hypothesis. This study confirmed that mechanical perturbation can impact xylem structural properties and result in altered plant water flow characteristics and cavitation resistance. Long‐term hydraulic acclimation in these herbaceous annuals was constrained by similar tradeoffs that constrain hydraulic properties across species.     

Mechanism of water‐stress induced cavitation in conifers: bordered pit structure and function support the hypothesis of seal capillary‐seeding

Resistance to water‐stress induced cavitation is an important indicator of drought tolerance in woody species and is known to be intimately linked to the anatomy of the xylem. However, the actual mechanical properties of the pit membrane are not well known and the exact mode of air‐seeding by which cavitation occurs is still uncertain. We examined the relationship between cavitation resistance and bordered pit structure and function in 40 coniferous species. Xylem pressure inducing 50% loss of hydraulic conductance (P₅₀, a proxy for cavitation resistance) varied widely among species, from ?2.9 to ?11.3 MPa. The valve effect of the pit membrane, measured as a function of margo flexibility and torus overlap, explained more variation in cavitation‐resistance than simple anatomical traits such as pit membrane, pit aperture or torus size. Highly cavitation resistant species exhibited both a high flexibility of the margo and a large overlap between the torus and the pit aperture, allowing the torus to tightly seal the pit aperture. Our results support the hypothesis of seal capillary‐seeding as the most likely mode of air‐seeding, and suggest that the adhesion of the torus to the pit border may be the main determinant of cavitation resistance in conifers.     

Photosynthetic pathway alters hydraulic structure and function in woody plants

Xylem structure and function is proposed to reflect an evolutionary balance between demands for efficient movement of water to the leaf canopy and resistance to cavitation during high xylem tension. Water use efficiency (WUE) affects this balance by altering the water cost of photosynthesis. Therefore species of greater WUE, such as C₄ plants, should have altered xylem properties. To evaluate this hypothesis, we assessed the hydraulic and anatomical properties of 19 C₃ and C₄ woody species from arid regions of the American west and central Asia. Specific conductivity of stem xylem (K_s ) was 16%–98% lower in the C₄ than C₃ shrubs from the American west. In the Asian species, the C₃ Nitraria schoberi had similar and Halimodendron halodendron higher K_s values compared with three C₄ species. Leaf specific conductivity (K_L ; hydraulic conductivity per leaf area) was 60%–98% lower in the C₄ than C₃ species, demonstrating that the presence of the C₄ pathway alters the relationship between leaf area and the ability of the xylem to transport water. C₄ species produced similar or smaller vessels than the C₃ shrubs except in Calligonum, and most C₄ shrubs exhibited higher wood densities than the C₃ species. Together, smaller conduit size and higher wood density indicate that in most cases, the C₄ shrubs exploited higher WUE by altering xylem structure to enhance safety from cavitation. In a minority of cases, the C₄ shrubs maintained similar xylem properties but enhanced the canopy area per branch. By establishing a link between C₄ photosynthesis and xylem structure, this study indicates that other phenomena that affect WUE, such as atmospheric CO₂ variation, may also affect the evolution of wood structure and function.     

Recovery of Populus tremuloides seedlings following severe drought causing total leaf mortality and extreme stem embolism

In contrast with other native Populus species in North America, Populus tremuloides (aspen) can successfully establish itself in drought‐prone areas, yet no comprehensive analysis has been performed on the ability of seedlings to withstand and recover from a severe drought resulting in complete leaf mortality. Here, we subjected 4‐month‐old aspen seedlings grown in two contrasting soil media to a progressive drought until total leaf mortality, followed by a rewatering cycle. Stomatal conductance (g_s), photosynthesis and transpiration followed a sigmoid decline with declining fraction of extractable soil water values. Cessation of leaf expansion occurred close to the end of the linear‐decrease phase, when g_s was reduced by 95%. Leaf mortality started after g_s reached the lowest values, which corresponded to a stem–xylem pressure potential (Ψ_xp) of ?2.0 MPa and a percent loss of stem hydraulic conductivity (PLC) of 50%. In plants with 50% leaf mortality, PLC values remained around 50%. Complete leaf mortality occurred at an average stem PLC of 90%, but all seedlings were able to resprout after 6–10 days of being rewatered. Plants decapitated at soil level before rewatering developed root suckers, while those left with a 4‐cm stump or with their stems intact resprouted exclusively from axillary buds. Resprouting was accompanied by recovery of stem hydraulic conductivity, with PLC values around 30%. The percentage of resprouted buds was negatively correlated with the stem %PLC. Thus, the recovery of stem hydraulic conductivity appears as an important factor in the resprouting capacity of aspen seedlings following a severe drought.     

Rootstock effects on xylem conduit dimensions and vulnerability to cavitation of <Emphasis Type="Italic">Olea europaea</Emphasis> L.

Two clones of Olea europaea L. were studied for their potential impact on hydraulic architecture and vulnerability to xylem cavitation, when used as rootstocks. The clones used were “Leccino Minerva” (LM), showing vigorous growth and “Leccino Dwarf” (LD) with strongly reduced growth. Self-rooted LM and LD plants as well as their grafting combinations were compared, namely, LM/LD (Leccino Minerva grafted onto Leccino Dwarf rootstock) and LD/LM (Leccino Dwarf grafted onto Leccino Minerva rootstocks). Plants with LD roots (LD and LM/LD) showed significantly reduced leaf surface area compared with plants with LM roots. Xylem conduits of LD shoots were 25% more numerous than in LM shoots. When grafted onto LM rootstocks, however, LD shoots produced consistently wider and longer vessels than measured in LD self-rooted plants. This caused LD/LM plants to increase stem vulnerability to cavitation with threshold pressures for cavitation (P _c) of less than 0.5 MPa compared with LD self-rooted plants that had P _c of over 2.0 MPa. By contrast, although LD rootstocks caused some reduction of vessel diameter and length of LM scions, their influence on LM hydraulic architecture was too small to reduce vulnerability to cavitation of LM scions with respect to that measured for LM self-rooted plants. Our conclusion is that although dwarfing rootstocks effectively reduce grafted plant size, they do not necessarily confer higher resistance to xylem cavitation to scions which would improve plant resistance to drought.     

Vulnerability to cavitation differs between current‐year and older xylem: non‐destructive observation with a compact magnetic resonance imaging system of two deciduous diffuse‐porous species

Development of xylem embolism during water stress in two diffuse‐porous hardwoods, Katsura (Cercidiphyllum japonicum) and Japanese white birch (Betula platyphylla var. japonica), was observed non‐destructively under a compact magnetic resonance imaging (MRI) system in addition to conventional quantitation of hydraulic vulnerability to cavitation from excised stem segments. Distribution of white and dark areas in MR images corresponded well to the distribution of water‐filled/embolized vessels observed by cryo‐scanning electron microscopy in both species. Water‐filled vessels were observed in MR images as white areas in Katsura and as white dots in Japanese white birch, respectively, and embolisms could be detected as a change to dark areas. The increase in the relative embolized area (REA: %) in the cross‐sectional area of total xylem during water stress, which was estimated from the binarized MR images, was consistent with the hydraulic vulnerability curves of these species. From the non‐destructive MRI observations, cavitation induced by water stress was shown to develop earlier in 1‐ or 2‐year‐old xylem than in the current‐year xylem in both species; that is, the vulnerability to cavitation differs between vessels in the current‐year xylem and those in older annual rings.     

The effects of intervessel pit characteristics on xylem hydraulic efficiency and photosynthesis in hemiepiphytic and non‐hemiepiphytic Ficus species

Xylem vulnerability to cavitation and hydraulic efficiency are directly linked to fine‐scale bordered pit features in water‐conducting cells of vascular plants. However, it is unclear how pit characteristics influence water transport and carbon economy in tropical species. The primary aim of this study was to evaluate functional implications of changes in pit characteristics for water relations and photosynthetic traits in tropical Ficus species with different growth forms (i.e. hemiepiphytic and non‐hemiepiphytic) grown under common conditions. Intervessel pit characteristics were measured using scanning electron microscopy in five hemiepiphytic and five non‐hemiepiphytic Ficus species to determine whether these traits were related to hydraulics, leaf photosynthesis, stomatal conductance and wood density. Ficus species varied greatly in intervessel pit structure, hydraulic conductivity and leaf physiology, and clear differences were observed between the two growth forms. The area and diameter of pit aperture were negatively correlated with sapwood‐specific hydraulic conductivity, mass‐based net assimilation rate, stomatal conductance (g_s), intercellular CO₂ concentration (C_i) and the petiole vessel lumen diameters (D_v), but positively correlated with wood density. Pit morphology was only negatively correlated with sapwood‐ and leaf‐specific hydraulic conductivity and D_v. Pit density was positively correlated with g_s, C_i and D_v, but negatively with intrinsic leaf water‐use efficiency. Pit and pit aperture shape were not significantly correlated with any of the physiological traits. These findings indicate a significant role of pit characteristics in xylem water transport, carbon assimilation and ecophysiological adaptation of Ficus species in tropical rain forests.     

Hydraulic architecture and water relations of Spartium junceum branches affected by a mycoplasm disease

Plant water relations, xylem anatomy and the hydraulic architecture of 1‐year‐old twigs of Spartium junceum, both healthy and affected by a phytoplasm disease, were studied. The disease causes twigs to be either shortened (witches broom disease, WBD) or flat (fasciate disease, FD). WBD twigs show a sevenfold increase in total leaf area, smaller and shorter xylem conduits, a higher stomatal conductance (g_l) and a decline of minimum leaf water potentials ( Ψ _l) below the turgor loss point. FD twigs had nearly twice the leaf area of the healthy controls as well as high g_l values and Ψ _l values below the turgor loss point. Moreover, significant differences between healthy and affected twigs in stem stomatal conductance (g_s) and in the total stem area were recorded. Affected twigs die back under drought stress, which is explained by a pronounced loss of hydraulic conductivity of the infected stems (40 and 60%) in FD and WBD as well as by the unfavourable ratio of weighted conduit radius ( Σ r⁴) to total surface area (A_t), so that the efficiency of the stem in supplying the whole transpiring area with water is strongly reduced.     

Water loss regulation to soil drought associated with xylem vulnerability to cavitation in temperate ring-porous and diffuse-porous tree seedlings

Key message

Sustainable stomatal opening despite xylem cavitation occurs in ring-porous species and stomatal closure prior to cavitation in diffuse-porous species during soil drought.

Abstract

To elucidate the relationship between water loss regulation and vulnerability to cavitation associated with xylem structure, stomatal conductance (g _s), defoliation, vulnerability curves, and vessel features were measured on seedlings of ring-porous Zelkova serrata and Melia azedarach, and diffuse-porous Betula platyphylla var. japonica, Cerasus jamasakura and Carpinus tschonoskii. Under prolonged drought conditions, the percentage loss of hydraulic conductivity (PLC) increased and g _s decreased gradually with decreasing predawn (Ψ_pd) or xylem water potential (Ψ_xylem) in Z. serrata. During the gentle increase of PLC in M. azedarach, g _s increased in the early stages of dehydration while leaves were partly shed. A sharp reduction in g _s was observed before the onset of an increase in the PLC for drying plants of the three diffuse-porous species, suggesting cavitation avoidance by stomatal regulation. In the ring-porous species, xylem-specific hydraulic conductivity (K _s) was higher, whereas the vessel multiple fractions, the ratio of the number of grouped vessels to total vessels, was lower than that in the diffuse-porous species, suggesting that many were distributed as solitary vessels. This may explain the gradual increase in the PLC with decreasing Ψ_xylem because isolated vessels provide less opportunity for air seeding. Different water loss regulation to soil drought was identified among the species, with potential mechanisms being sustainable gas exchange at the expense of xylem dysfunction or partial leaf shedding, and the avoidance of xylem cavitation by strict stomatal regulation. These were linked to vulnerability to cavitation that appears to be governed by xylem structural properties.